The Neomiodontids: Brackish-Water Bivalves of the Mesozoic

Cast from internal mould of Myrene tetoriensis, from here.

During the early part of the Cretaceous, an area corresponding to the modern Sea of Japan was occupied by a massive brackish-water lake that has been called Lake Tetori, after the geological Tetori Formation that it left behind it. My impression is that Lake Tetori was not an overly hospitable place: warm, shallow, and probably low in oxygen, it was home to a fairly depauperate fauna dominated by only two species of clam, known as Tetoria yokoyamai and Myrene tetoriensis (Kondo et al. 2006). Tetoria I shall leave for another time; Myrene (and its ilk) is the one I want to look at today.

Myrene tetoriensis belonged to a now-extinct family of bivalves known as the Neomiodontidae that lived during the Jurassic and Cretaceous periods. Neomiodontids were most diverse in the northern continents, though species have also been assigned to this family from India and Australia (Moore 1969). Not dissimilar in appearance to a modern pipi, though generally smaller in size, neomiodontids are primarily known from brackish-water or freshwater deposits. The late Triassic/early Jurassic saw something of a flush of bivalve lineages in low salinity environments: the separation of the ex-Pangaean continents resulted in an increase in continental margins, while high carbon dioxide concentrations in the atmosphere stymied the growth of calcium-heavy marine forms (Kondo & Sano 2009).

A rock full of fossils of Neomiodon, from here.

Most neomiodontids were found in sandy habitats, though Myrene tetoriensis lived in mud. They were shallow burrowers, living buried in the sand with the tip of the shell at surface level. Deep-burrowing bivalves possess elongate tubular siphons through which they breath and feed; the mantle boundary inside the shell of such species has a cavity called the pallial sinus into which the siphons can be retracted. In neomiodontids, the pallial sinus is undeveloped, indicating a proportional lack of development of any siphons. Neomiodontids would have mostly been suspension feeders, capturing food particles floating in the water; because of its muddier habitat, Myrene may have been a deposit feeder (Nishida et al. 2013). The narrow, relatively slim shape of neomiodontid shells suggests that they could probably burrow into their substrate rapidly if they became exposed by water action or potential predators.

When the Neomiodontidae was first established as a distinct family, Casey (1955) suggested that it could include the ancestors of the Sphaeriidae, a living group of small freshwater clams known as the pea clams. If this was the case, then neomiodontids did not truly go extinct during the Cretaceous but live on in their descendants. However, more recent authors do not seem to support this relationship. Kondo et al. (2006) note that the decline of brackish-water shallow burrowers such as neomiodontids correlated with the diversification of deeper-burrowing families and suggest a causal connection between the two, but it is worth noting that Tetoria, mentioned above as peaceful cohabitant of Myrene, was a deep-burrower. Nishida et al. (2013) see the extinction of neomiodontids somewhat differently. Citing the often transient nature of the habitats preferred by neomiodontids and other non-marine bivalves, they suggest that the neomiodontids were not a single lineage but represented numerous independent colonisations of non-marine habitats by members of the related marine family Arcticidae, with the 'neomiodontid' habitus the result of convergent evolution.In this view, the reasons for 'neomiodontid' extinction should be sought not with the neomiodontids themselves, but with the extinction of their arcticid progenitors.

REFERENCES

Casey, R. 1955. The Neomiodontidae, a new family of the Arcticacea (Pelecypoda). Proceedings of the Malacological Society 31: 208–222.

Kondo, Y., T. Kozai, N. Kikuchi & K. Sugawara. 2006. Ecologic and taxonomic diversification in the Mesozoic brackish-water bivalve faunas in Japan, with emphasis on infaunalization of heterodonts. Gondwana Research 10: 316–327.

Kondo, Y., & S. Sano. 2009. Origination of extant heteroconch families: ecological and environmental patterns in post-Paleozoic bivalve diversification. Paleontological Research 13 (1): 39–44.

Moore, R. C. (ed.) 1969. Treatise on Invertebrate Paleontology pt N. Mollusca 6. Bivalvia vol. 2. The Geological Society of America, Inc., and The University of Kansas.

Nishida, N., A. Shirai, K. Koarai, K. Nakada & M. Matsukawa. 2013. Paleoecology and evolution of Jurassic–Cretaceous corbiculoids from Japan. Palaeogeography, Palaeoclimatology, Palaeoecology 369: 239–252.

Juniper Gall Midges

Sometimes, the evidence of an insect's presence may be much more visible than the insect itself. Imagine passing by a common juniper tree Juniperus communis and seeing a structure like the one in the photo above (copyright Jean-Yves Baugnée). You might think it was some sort of reproductive structure. You would be right, though it is not the tree that is reproducing. This is the gall of a juniper gall midge Oligotrophus juniperinus, and were you to cut the gall in half you would possibly find a single gall midge larva lurking within. Many insects (and other animals) cause the development of galls on their host plants, thus providing themselves with both shelter and food in one convenient location.

Male Oligotrophus betheli, from Simova-Tošić et al. (2010).

The adult gall midge is a minute, very delicate fly, unlikely to be spotted by the casual observer. Gall midges are classified in the family Cecidomyiidae, an extremely diverse group of which not all members cause galls as larvae (some feed on plants without causing galls, others feed on fungi, a few are even predators or parasitoids). Cecidomyiids are divided between a number of subfamilies and tribes, with Oligotrophus belonging to the tribe Oligotrophini. In the past, this tribe has been used to cover a heterogeneous mix of relatively unspecialised cecidomyiids, but the most recent classification of the tribe strips it down to two genera, Oligotrophus and Walshomyia, found in the Holarctic region (Harris et al. 2006). Adults of these genera have legs with simple tarsal claws and long empodia (the soft pads between the claws), and as larvae they all live in galls on trees of the cypress family Cupressaceae. The exact form of the gall produced may differ between species, and it is often (though not always) possible to determine the species responsible for a gall by its form. For instance, three species that cause galls on Juniperus communis in Europe are Oligotrophus juniperinus, O. panteli and O. gemmarum. The first two species have galls formed from whorls of leaves pressed into a vase shape, but whereas in galls of O. juniperinus the leaves splay outwards towards the tip, in galls of O. panteli they remain parallel. The third species, O. gemmarum, has much smaller galls formed from only slightly modified buds; though very different from mature galls of the other two species, they may be confused with young undeveloped galls (Harris et al. 2006).

Despite their diversity, and the fact that some species are economically significant to humans, cecidomyiids are not a widely studied group. Part of the reason for this is that their small size and build makes them difficult to handle; diagnostic work on adults often requires slide-mounting them. I have made one not-very-successful attempt at slide-mounting cecidomyiids, and I can confirm that it is a fiddly process. Because the different body parts often have to be examined from different angles, slide-mounting first requires dissection of the animal into sections (so, for instance, the head can be placed on the slide face-on, the body side-on, and the terminalia top-up). In my experience, instructions for slide-mounting animals requiring such dissections will always tell you to arrange the various bits appropriately on the cover-slip before placing the slide (or the other way around, if you prefer). And if you know how to attach slide to cover-slip without having all your carefully arranged body parts immediately zooming off to a completely different spot on the slide from where you put them, then you're a far more skillful slide preparer than I am.

REFERENCES

Harris, K. M., S. Sato, N. Uechi & J. Yukawa. 2006. Redefinition of Oligotrophus (Diptera: Cecidomyiidae) based on morphological and molecular attributes of species from galls on Juniperus (Cupressaceae) in Britain and Japan. Entomological Science 9: 411–421.

Simova-Tošić, D., D. Graora, R. Spasić & D. Smiljanić. 2010. Oligotrophus betheli Felt (Diptera: Cecidomyiidae), a new species in the fauna of Europe. Arch. Biol. Sci. 62 (4): 1219–1221.

Mites in Red Velvet

Adult Platytrombidium fasciatum, copyright Walter Pfliegler.

Mites in red velvet,
decorated with stripes.
Completing their diet,
hunting down eggs from flies*.

*With apologies to Justin Hayward.

Among the mites most likely to be seen by the casual observer are the various species of active predators known as red velvet mites. They grow to relatively large sizes for a mite (the species in the photo above can get up to 2.5 mm long), they are brightly coloured and they can often be seen moving about in search of food. As well as the colour, the name 'red velvet mite' refers to their dense covering of setae giving them almost a teddy-bearish appearance. There aren't many mites that could be described as cuddly, but these are arguably among them (at least as adults, as explained below).

Red velvet mites form a number of families in the mite clade Parasitengonina. Earlier posts on this site (here and here) have already described the somewhat complicated life cycles of parasitengonines, but to recap briefly: parasitengonines start their lives as parasitic larvae, followed by a dormant 'pupa-like' stage, followed by an active predatory nymph, then another dormant 'pupa', and finally the active predatory adult. Whereas differences between the active nymphs and adults are slight (kind of raising the question as to why the intervening dormant phase), differences between adults and larvae are significant. From their appearance alone, there is no way of telling whether a given larval form corresponds to a given adult, and connecting the two requires challenging indirect methods such as brood-rearing. Nevertheless, both forms are commonly encountered: not only are adults significant micro-predators, the larvae are often found attachned to insects and other arthropods. Some larval species, commonly known as chiggers, attack vertebrates such as humans and so are even more well-studied. Because of the resulting need to classify both adults and larvae without an easy way to connect the two, a kind of double taxonomy has developed with many parasitengonines. Adults and larvae are treated as if they were separate 'genera' and 'species', with separate names for each. Sometimes a larval 'species' may be successfully connected to an adult 'species' and the two can be synonymised, but many taxa remain that are known only from one or the other.

The genus Platytrombidium, belonging to the velvet mite family Microtrombidiidae, was established in 1936 on the basis of adults, but its larval form was not described until 2005. A number of species have been assigned to this genus from various parts of the world but, as a result of obtaining better descriptions of both adult and larva, Gabryś et al. (2005) restricted it to three species known from the Palaearctic region (Europe and northern Asia). Adult Platytrombidium are characterised by an even covering of stout, uniform setae covered with delicate setules; when alive, they are even more readily recognised from their transverse white stripes across the body. As adults and active nymphs, Platytrombidium fasciatum (the best-known species in the genus and the only one with known larvae) feed on fly eggs. Their larvae are also parasites on drosophilids and similar small flies, most often found attached to the dorsal surface of the abdomen (Gabryś et al. recorded one larva found attached to its host's eye).

Most of the confusion about the taxonomy of Platytrombidium has revolved around its relationship to the very similar genus Atractothrombium. For a long time, the only recognised difference between the two was whether the setae on the body were pointed (Platytrombidium) or blunt (Atractothrombium). Needless to say, this was not a very clear character, and might appear to vary even over the surface of a single individual. Nevertheless, Gabryś et al. (2005) found that they were able to distinguish the type species of the two genera by features of the adult palps and larval claws (Atractothrombium sylvaticum is also evenly dark red, lacking the white stripes of Platytrombidium fasciatum). They also differ in habits: both are predators and parasites of flies but whereas P. fasciatum is found in drier habitats such as gardens and parks, A. sylvaticum prefers damp habitats that flood regularly, such as reed beds and salt marshes.

REFERENCES

Gabryś, G., A. Wohltmann & J. Mąkol. 2005. A redescription of Platytrombidium fasciatum (C. L. Koch, 1836) and Atractothrombium sylvaticum (C. L. Koch, 1835) (Acari: Parasitengona: Microtrombidiidae) with notes on synonymy, biology and life cycle. Annales Zoologici 55 (3): 477–496.

Stars and Blessings

Yellow starthistle Centaurea solstitialis, copyright Franco Folini.

The first thing that struck me when I was looking up material on Centaurea was how evocative some of the vernacular names associated with this genus are: starthistle, blessed thistle, dusty miller, sweet sultan. Centaurea, the starthistles and knapweeds, is a genus of composite-flowered plants native to Eurasia and northern Africa, with the highest diversity of species in the Mediterranean region. A handful of species have been spread to other parts of the world in association with humans; a handful of these are significant pasture pests such as spotted knapweed C. maculosa and yellow starthistle C. solstitialis, whereas others such as dusty miller C. cineraria are grown as garden plants. Centaurea is a large genus: depending on how you count them, it may contain anywhere between 300 and 700 species. The greater number of species are perennial herbs, but the genus varies from small spiny shrubs to low spreading annuals (Wagenitz 1986). Some arise from a single central tap-root; others grow from spreading rhizomes. Some species have spiny leaves and conform to our general idea of a 'thistle'; others do not. The leaves are often deeply divided at the base of the plant, becoming entire towards the top. Flowerheads may be borne singly or in a corymbiform arrangement (a flat-topped cluster); the phyllaries (the bracts surrounding the flowerhead) often extend outwards around the head, and may be themselves tipped with spines.

Squarrose knapweed Centaurea triumfettii, copyright Kristian Peters.

With a genus of this size, it should be hardly surprising that taxonomic complications are involved. Long recognised as morphologically diverse, it has been confirmed as polyphyletic by more recent molecular analyses (Garcia-Jacas et al. 2001). The majority of Centaurea species fall within a single derived clade within the composite subtribe Centaureinae, united both by molecular data and by a number of morphological synapomorphies including adaptations for myrmecochory, dispersal of the seeds by ants (the seeds carry an attached oily body called an elaiosome; ants carry the seeds back to their nest where they may eat the elaiosome but leave the seed to sprout). A handful of species, though, lack these synapomorphies and lie in scattered segregate clades among the remainder of the Centaureinae. Some of these segregate clades, such as the former section Psephellus, have been straightforwardly promoted to the status of separate genera. One small segregate clade, however, is a little more problematic because it happens to include the north African Centaurea centaurium, the original type species of the genus Centaurea. Under normal circumstances, then (other than lumping the entirety of centaureines in a single genus), the name Centaurea would apply only to this small clade (including only about a dozen species) while the hundreds of species in the main 'Centaurea' clade would have to be renamed. In this case, the name with priority for this large clade would be Cnicus, generally used to date for only a single species, the blessed thistle Cnicus benedictus (no, I haven't been able to establish why it is called the 'blessed thistle'; I have found references to a tradition of medicinal use for this species, including its supposedly encouraging milk production in nursing mothers, but I haven't been able to confirm if this is the reason for the name). In order to stave off this nomenclatural turmoil, it has been proposed that the official type species of Centaurea be changed to a member of the main clade (Greuter et al. 2001), so this clade keeps the name Centaurea (and the blessed thistle becomes referred to as Centaurea benedicta) whereas the small clade including the prior type species becomes known as the genus Rhaponticoides. I haven't found whether a final decision has been made on this proposal (the process for such nomenclatural decisions is a bit more involved for plants than animals, requiring an open vote at an international botanical conference rather than just being decided on directly by a select committee) but it seems to have general support. Less certain is the status of the cornflowers of the section Cyanus, which some have proposed recognising as a separate genus but which is closely related to the main clade, making the case for its separation a bit less compelling.

REFERENCES

Garcia-Jacas, N., A. Susanna, T. Garnatje & R. Vilatersana. 2001. Generic delimitation and phylogeny of the subtribe Centaureinae (Asteraceae): a combined nuclear and chloroplast DNA analysis. Annals of Botany 87: 503–515.

Greuter, W., G. Wagenitz, M. Agababian & F. H. Hellwig. 2001. (1509) Proposal to conserve the name Centaurea (Compositae) with a conserved type. Taxon 50: 1201–1205.

Wagenitz, G. 1986. Centaurea in south-west Asia: patterns of distribution and diversity. Proceedings of the Royal Society of Edinburgh, Section B, Biological Sciences 89: 11–21.

The Ostrich: From Whence this Derpy Horror?

Male and two female ostriches Struthio camelus, copyright Yathin S. Krishnappa.

Ostriches are widely known for two things: firstly, that they are the largest living bird by a quite respectable margin, and secondly, that they look ridiculous. Seriously, is there anyone out there who can look at the animals in the picture above and not think them ludicrous. Though I am, admittedly, invoking the luxury of distance: my uncle spent a year or two raising ostriches back during the brief boom of ostrich farming in New Zealand in the early 2000s, and I can say from experience that what looks humorous from afar is, close up, intimidating in a way no other animal is. They're just so tall*.

*Not to mention their well-earned reputation for gobbling down any item that attracts their attention. There are numerous stories out there demonstrating that wearing jewellery in an ostrich enclosure is a bad idea.

The modern ostrich is commonly regarded as a single species, Struthio camelus, found in savannah and semi-desert habits around Africa. There are some grounds for recognising the Somali ostrich S. molybdophanes of the Horn of Africa as a separate species—it is both genetically and morphologically divergent from other ostrich populations (for instance, its skin is blue rather than pink or red), and there is a small amount of overlap in range between Somali and typical ostriches—but this question remains open. Other subspecies are the North African ostrich S. c. camelus, the southern ostrich S. c. australis, and the Masai ostrich S. c. massaicus of Tanzania and Kenya. A fifth subspecies, the Arabian ostrich S. c. syriacus, became extinct around 1940, though it is worth noting that mitochondrial DNA extracted from specimens of Arabian ostriches in the British Museum did not separate them from the North African ostrich (Robinson & Matthee 1999). Ostriches can not really be confused with any other modern bird: not only is their remarkable size (matched by the remarkable size of their eggs), but they are the only birds to have reduced the number of toes to just two, with only the third and fourth toes of the standard bird foot remaining. This feature is generally presumed to be related to their cursorial lifestyle.

More evidence that ostriches are just daft. Copyright Georges Olioso.

Ostriches are members of the group of birds known as palaeognaths, that also includes such birds as the emu, kiwis, cassowaries, rheas and tinamous (the flightless members of the palaeognaths are commonly referred to as the ratites, but recent studies have cast doubt on whether flightlessness in the palaeognaths has a single origin). Phylogenetic relationships within the palaeognaths have been shuffled about considerably over the years (and even now are probably not really settled), but it is generally agreed that ostriches probably diverged from their nearest living relatives a long time ago (Burleigh et al. [2015], for instance, place them as the sister taxon to all other palaeognaths). Just how long ago we can't really say, the early fossil record of ostriches (and palaeognaths in general) being pretty dire. The heron-sized middle Eocene Palaeotis weigelti from Germany has been suggested to be a direct relative of ostriches but the evidence for this is equivocable (Mayr 2009). The earliest undoubted ostrich is the early Miocene Struthio coppensi from Namibia, and this is already similar enough to modern ostriches to be placed in the same genus.

Fossil ostriches are known from southeastern Europe to China, and survived across much of Asia until the Pleistocene. Several species have been named, but the usual vagaries of preservation make it debatable how many are distinct. Matters are complicated by several 'species', such as the Ukrainian Struthio chersonensis, that have been named based on fossil eggshells, raising questions as to whether such names can or should be applied to associated body fossils. Also unknown are the phylogenetic relationships between modern and fossil ostriches: whether the Eurasian ostriches represented a single or multiple dispersals out of Africa, or even whether ostriches may have originated in Eurasia*.

*It was suggested at one point that ostriches may have originally come from India, only dispersing to Africa after the subcontinent latched onto the rest of Eurasia. Support for this was based on the phylogenetic hypotheses that ostriches and the South American rheas formed an exclusive clade within the palaeognaths, and that the divergence of the flightless ratites was directly influenced by the division of the Gondwanan supercontinent (a South American-African connection being inconsistent with Africa being the first part of Gondwana to be separated). As support for both these arguments has declined, the need to somehow get ostriches out of Africa has evaporated.

The earliest known ostrich, the aforementioned Struthio coppensi, was a smaller and more slender bird than the modern ostrich, but some fossil species were larger. Perhaps the tallest ostrich species was S. oldowayi of the Tanzanian Pleistocene, which had a femur about a third as long again as the modern species. The femur of the Georgian Plio-Pleistocene S. dmanisiensis was not quite as long as that of S. oldowayi but it was considerably more robust, suggesting a proportionally solidly-built bird (Vekua 2013). Struthio brachydactylus (which sometimes moonlights as S. chersonensis) from the Miocene of Ukraine was also robustly built, albeit probably no taller (if not shorter) than a modern ostrich, but its main distinction lies in it taking the toe reduction seen in other ostriches even further. The fourth toe was reduced, with more weight placed on the third toe, making this species functionally almost single-toed (Boev & Spassov 2009).

REFERENCES

Boev, Z., & N. Spassov. 2009. First record of ostriches (Aves, Struthioniformes, Struthionidae) from the late Miocene of Bulgaria with taxonomic and zoogeographic discussion. Geodiversitas 31 (3): 493–507.

Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

Mayr, G. 2009. Paleogene Fossil Birds. Springer.

Robinson, T. J., & C. A. Matthee. 1999. Molecular genetic relationships of the extinct ostrich, Struthio camelus syriacus: consequences for ostrich introductions into Saudi Arabia. Animal Conservation 2: 165–171.

Vekua, A. 2013. Giant ostrich in Dmanisi fauna. Bulletin of the Georgian National Academy of Sciences 7 (2): 143–148.

Syntomodrillia

Syntomodrillia cybele, copyright Korina Sangiouloglou.

Time, I think, for another visit to the often overlooked hotbed of gastropod diversity that is the 'turrids'. As alluded to here and here, these are the less differentiated members of the cone shell superfamily Conoidea, treated in the past as a single family Turridae but now classified into several different families.

Syntomodrillia is a genus in the conoid family Drilliidae. These are small shells, with recent species no more than a centimetre in length (Woodring 1970). Recent species of Syntomodrillia are found only in the American tropics, mostly in the Caribbean and the Gulf of Mexico, with a single species S. cybele (the one shown above) at the Galapagos Islands. The fossil record, however, may indicate a broader range for Syntomodrillia in the past, as Powell (1966) assigned species to this genus from the Oligocene to the Pliocene of Australasia and Okinawa. Syntomodrillia is similar in appearance to another drilliid genus, the somewhat magnificently named Splendrillia, and has been treated by some authors as a subgenus of the latter. Among the features distinguishing the two is the appearance of the longitudinal ribs running down the shell: in Syntomodrillia, the ribs completely cross each whorl, but in Splendrillia they are interrupted on the shoulder. The protoconch (larval shell) also differs between the two, with that of Syntomodrillia being slender with two whorls, whereas that of Splendrillia is broadly rounded and paucispiral (Powell 1966). This may indicate that the larval stage of Syntomodrillia is slightly longer and/or more active than that of Splendrillia.

Radula of Splendrillia, from Kantor & Puillandre (2012); mt = marginal teeth.

As described in an earlier post, the Conoidea have alternatively been known as the 'Toxoglossa' because many conoids have the radula modified for the injection of toxins (taken to the utmost in the cone shells, which may be capable of killing humans). The median and lateral teeth of the radula are reduced or lost, and the marginal teeth turn into disposable syringes. The Drilliidae, however, have not gone down this path: they retain a radula with well-developed saw-like lateral teeth. Though records of drilliid diet are decidedly sparse, they probably hunt soft-bodied prey by actively grabbing and tearing it, in contrast to the more refined eating habits of other conoids.

REFERENCES
Kantor, Y. I., & N. Puillandre. 2012. Evolution of the radular apparatus in Conoidea (Gastropoda: Neogastropoda) as inferred from a molecular phylogeny. Malacologia 55 (1): 55–90.

Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae: an evalution of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Auckland Institute and Museum 5: 1–184, 23 pls.

Woodring, W. P. 1970. Geology and paleontology of Canal Zone and adjoining parts of Panama: description of Tertiary mollusks (gastropods: Eulimidae, Marginellidae to Helminthoglyptidae). Geological Survey Professional Paper 306-D: 299–452, pls 48–66.

Small Carrion Beetles: A Bunch of SBBs

A fairly typical small carrion beetle, Catops tristis, copyright Trevor and Dilys Pendleton.

Anyone who takes on the task of beetle identification will soon discover that (to agree with Haldane) their sheer diversity can be overwhelming. Bird-watchers often complain about the challenges of identifying what they refer to as LBJs, Little Brown Jobs, but entomologists may have as much if not more to complain about when faced with the prospect of SBBs: Small Brown Beetles. The features marking a particular SBB as one family or another are often (at least to a novice) difficult to distinguish; members of unrelated families may look remarkably similar, whereas close allies may look surprisingly different.

The Leiodidae are, for the most part, firmly in the ranks of SBBs. This taxonomically small but morphologically diverse family is hard to come up with a coherent description for: though modern coleopterists have little doubt that they form a coherent clade, certain subgroups have become notably divergent. At least one leiodid, the beaver parasite Platypsyllus castoris, barely even looks like a beetle at all and was classified for a brief period in the 1800s as a distinct order of insects. Nevertheless, most leiodids are recognised by the structure of their antennae: the five-segmented club at its end has a distinct constriction as the eight antennal segment is smaller than the seventh and ninth segments on either side. Many leiodids are scavengers of plant or animal matter, but some are fungivores and a few (as already indicated) are parasites of mammals.

Small carrion beetles of the genus Sciodrepoides feeding on a deer carcass; copyright Stephen Cresswell.

Among the various subgroups of the Leiodidae are the Cholevini, commonly known as small carrion beetles. As their name indicates, these mostly feed on the remains of dead animals, though at least some are not above scavenging on other decaying matter. Some species are found in subterranean habitats, such as caves or the burrows of rodents, feeding on guano or other refuse. The Cholevini are one of the tribes in the leiodid subfamily Cholevinae, which has sometimes been treated in the past as a separate family Cholevidae or Catopidae. The Cholevinae differ from most other leiodids in the presence of an occipital carina or crest on the back of the head; such a carina is also present in the parasitic Leptininae, which Peck (1990) speculated to be derived from the cholevines. Members of the tribe Cholevini differ from other Cholevinae in having the setae on the elytra irregularly arranged (vs arranged in rows), giving the elytra a granular rather than a striate appearance.

Members of the Cholevini are mostly found in the Holarctic region, with only a few species in the Oriental region and none further south (Peck & Cook 2002). The greatest diversity in the group is found in Eurasia; only four of the 24 genera are found in North America, and only one of these (the monotypic Catoptrichus frankenhauseri) is unique to that continent. For the most part, cholevins do not vary much in appearance, and species are difficult to distinguish without examining the genitalia (these are true SBBs). Catoptrichus frankenhauseri has distinctive antennae, with lateral projections on either side of each segment(C. frankenhauseri is also noteworthy for the manner of its initial discovery, with the type specimen being collected from a human cadaver [Peck & Cook 2002]). Some of the subterranean species of cholevins have reduced eyes or wings, and a handful of species are entirely flightless.

REFERENCES

Peck, S. B. 1990. Insecta: Coleoptera Silphidae and the associated families Agyrtidae and Leiodidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1113–1136. John Wiley & Sons.

Peck, S. B., & J. Cook. 2002. Systematics, distributions, and bionomics of the small carrion beetles (Coleoptera: Leiodidae: Cholevinae: Cholevini) of North America. Canadian Entomologist 134: 723–787.